Research Article |
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Corresponding author: Krittiya Trivalairat ( krittiya.chn@mahidol.ac.th ) Corresponding author: Benchapol Lorsunyaluck ( benchapol.lo@wu.ac.th ) Academic editor: Minh Duc Le
© 2026 Poramad Trivalairat, Krittiya Trivalairat, Nawee Moungkhaek, Simon Caponov, Benchapol Lorsunyaluck.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Trivalairat P, Trivalairat K, Moungkhaek N, Caponov S, Lorsunyaluck B (2026) Acanthosaura syrax sp. nov., a new species of mountain horned dragon from high-elevation habitats of western Thailand (Squamata, Agamidae). ZooKeys 1287: 47-76. https://doi.org/10.3897/zookeys.1287.186636
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A new species of mountain horned dragon, Acanthosaura syrax sp. nov., is described from the high–elevation evergreen forests of Mae Wong National Park in western Thailand. Morphological comparisons distinguish the new taxon from its congeners by its smaller body size and the fact that it possesses the smallest nuchal and dorsal scales recorded within the genus. Phylogenetic analysis of mitochondrial ND2 and COI gene sequences further supports this distinction, identifying the new species as a reciprocal monophyletic lineage with 76% and 99% nodal support in maximum–likelihood analyses and 0.78 and 1.00 nodal support in Bayesian Inference analyses based on ND2 and COI genes, respectively. Interspecific pairwise genetic distances between A. syrax sp. nov. and other congeners range from 14.1–15.9% in ND2 gene and 14.96–28.13% in COI gene, while intraspecific variation remains low at 0–0.01% and 0–0.34% in ND2 and COI, respectively. This discovery brings the number of Acanthosaura species to 23 and highlights the significant and unique biodiversity found within the high–altitude evergreen ecosystems of western Thailand.
COI, Mae Wong National Park, morphology, ND2, reptile, Tak–Kamphaeng Phet Province, taxonomy, tropical rainforest
Acanthosaura Gray, 1831, commonly known as mountain horned dragons or pricklenape lizards, is a genus of agamid lizards characterized by a distinctive row of nuchal and dorsal crests (
In Thailand, the known diversity of Acanthosaura taxa has increased in recent years, currently comprising at least seven species: Acanthosaura armata (Gray, 1827) from the southern region; A. aurantiacrista Trivalairat, Kunya, Chanhome, Sumontha, Vasaruchapong, Chomngam & Chiangkul, 2020 from the north-western region; A. cardamomensis Wood, Grismer, Grismer, Neang, Chav & Holden, 2010 from the eastern region; A. crucigera Boulenger, 1885 from the western region; A. lepidogaster (Cuvier, 1829) from the northern region; A. phuketensis Pauwels, Sumontha, Kunya, Nitikul, Samphanthamit, Wood & Grismer, 2015 from Phuket Island and the south-western region; A. meridiona Trivalairat, Sumontha, Kunya & Trivalairat, 2022 from the southern region; and A. rubrilabris Liu, Rao, Hou, Orlov, Ananjeva & Zhang, 2022 from the northern region (
Mae Wong National Park, Tak–Kamphaeng Phet Province, Thailand is an area characterized by high-elevation evergreen forests reaching up to 1,964 m a.s.l., with a cold temperate climate year-round (
Specimens were conducted within range of Mae Wong National Park, Tak–Kamphaeng Phet Province, Thailand (16°06'01.6"N, 99°06'28.0"E) by hand. Photographs were taken to document the coloration pattern in life prior to euthanasia. Euthanasia was performed using a freezing technique at –10 °C for one week, at this species can hibernate in low-temperature conditions for a few days. The specimens were fixed in 10% neutral buffered formalin (NBF) for one week to preserve tissues. Following fixation, they were rinsed with distilled water and transferred to 70% ethanol for storage.
For molecular analyses, fresh liver samples were dissected into small pieces and stored in absolute ethanol. All specimens were deposited at Natural History Museum, National Science Museum, Technopolis, Pathum Thani Province, Thailand (THNHM).
Meristic and measured morphological characters were noted on the left side followed
Molecular data were generated from specimens collected in the Mae Wong National Park Range. To establish the phylogenetic context, these data were compared against sequences for various Acanthosaura taxa and a designated outgroup, Calotes emma Gray, 1845, which were retrieved from the GenBank database Appendix
A 698-base pair (bp) fragment of the NADH dehydrogenase subunit 2 (ND2) gene and 600–bp fragment of the Cytochrome c oxidase subunit I (COI) gene were amplified via Polymerase Chain Reaction (PCR) using EP0402 TAQ DNA POLYMERASE. The following primer pair was used for amplification: METF6 (L4437a; 5’–AAG CTT TCG GGC CCA TAC C–3’) and ACANTHND2.833. R1 (5’–AGG GAG GTT ATT GTT GCT AG–3’) for ND2; Chmf4 (5’–TYT CWA CWA AYC AYA AAG AYA TCG G–3’) and Chmr4 (5’–ACY TCR GGR TGR CCR AAR AAT CA–3’) for COI (
The thermal cycling protocol for the ND2 gene amplification was performed for 32 cycles as follows (
The DNA sequences were managed and prepared for phylogenetic analysis. Sequence editing and alignment were performed using the Clustal Omega (accessible at https://www.ebi.ac.uk/Tools/msa/clustalo/), implemented through the MEGA6 software suite (
The phylogenetic reconstructions were conducted using two complementary approaches: Maximum Likelihood (ML) and Bayesian Inference (BI). The optimal models of molecular evolution for both datasets were selected using ModelFinder (
The ML analysis was performed using IQ-TREE v. 2.2.2.6 (
Clade support was considered statistically significant when ultrafast bootstrap (UFB) values were ≥ 95% (
Acanthosaura armata (Gray, 1827).
Body somewhat compressed, usually with a rather sharp dorsal ridge and a dorsal crest larger in males; dorsal scales unequal, heterogenous; a gular pouch may be present; a fold cross shoulder usually conspicuous; tympanum naked or partially covered by scales; tail somewhat flattened, the scales keeled and larger than dorsal body scales, longer than broad; no femoral or preanal pores (
Acanthosaura species can be found across Southeast Asia and extending southward to Malay Peninsula (Table
| Country | Acanthosaura species | Reference |
|---|---|---|
| Cambodia | A. capra, A. cardamomensis, A. coronata, A. crucigera; |
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| China (southern region) | A. armata, A. lepidogaster, A. liui, A. longicaudata, A. rubrilabris, A. tongbiguanensis | Gray (1827); |
| Laos | A. coronata, A. lepidogaster, A. nataliae, A. prasina |
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| Malay Peninsula | A. armata, A. bintangensis, A. titiwangsaensis | Gray (1827); |
| Myanmar | A. armata, A. crucigera, A. lepidogaster | Gray (1827); |
| Thailand | A. armata, A. aurantiacrista, A. cardamomensis, A. crucigera, A. lepidogaster, A. meridiona, A. phuketensis, A. rubrilabris | Gray (1827); |
| Vietnam | A. brachypoda, A. capra, A. cardamomensis, A. coronata, A. cuongi, A. grismeri, A. lepidogaster, A. murphyi, A. nataliae, A. phongdienensis, A. prasina |
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Holotype: Thailand • 1 mature male; Kamphaeng Phet Province, Mae Wong District, Mae Wong National Park, Khun Nam Yen; 16°05'43.5"N, 99°07'25.2"E; 1,340 m a.s.l.; 23 Apr 2025; collected by Poramad Trivalairat & Krittiya Trivalairat; THNHM31129 (Fig.
A small-sized species (maximum SVL 101.2 mm for females and 69.6 mm for males) with a single tiny conical spine above the posterior margin of the eye (maximum PS 2.7 mm for females and 2.0 mm for males); small spine on the occiput between the tympanum and the nuchal crest (maximum OS 4.1 mm for females and 2.4 mm for males); tympanum scaleless, small, roundish; less developed gular pouch; present small scales intermixed with three set of medium keeled scales line transversely from dorsal to lateral on the flanks; nuchal crest with six or seven tiny, keeled, compressed, asymmetrical triangular spines ascending from the first to the fifth or sixth, then decreasing in height; narrow diastema of 5–8 scales between the nuchal and vertebral crests; vertebral crest composed of small sized saw-like triangular scales beginning in the shoulder region and decreasing in size until the base of the tail; tail 1.06–1.16 of SVL (85.5–106.9 mm for females and 72.4–77.6 mm for males); black collar patch present; black eye patch absent.
Adult male. SVL 67.0 mm; TL 77.6 mm, tail complete; HL 21.1 mm; head length is one-third the length of the body (HL/SVL 0.32), narrow (HW/SVL 0.21), moderately tall (HD/HL 0.53), triangular in dorsal and lateral views; snout moderately short (SL/HL 0.33); rostrum wide (RW/RH 2.33), steeply sloping anteriorly; canthus rostralis prominent, forming a large projecting shelf extending above the eye, composed of 12 large scales; shelf terminates with a notch anterior to the postorbital spine; rostrum wide in size, rectangular, bordered laterally by the first supra labials (totally two scales from left and right) and posteriorly by four upper smaller scales; nasal roundish, surrounded by two prenasal anteriorly, three postnasals posteriorly and one subnasal; six scales between the nasal scales; elongate supra nasal scales; moderate scales above the orbit weakly keeled; three rows of slightly keeled scales below the orbit extending from the posterior margin of the nasal to post orbits; eye moderately large (EYE/HL 0.21), orbit moderately large (ORBIT/HL 0.35); prefrontal and frontal slightly keeled and smaller than the scales between the orbit and supralabials occipital scales weakly keeled; large parietal; tiny conical epidermal spine (1.30 mm) above the posterior margin of the eye, posteriorly pointed, surround by four small lanceolate scales; suborbital scales small, slightly keeled, extending in a row of five equal large scales from below the posterior margin of the eye to the anterior margin of the tympanum, decreasing in size posteriorly; tiny conical epidermal spine (1.80 mm) larger than the postorbital spine, posteriorly pointing backward, surrounded by a rosette of five small lanceolate scales; tympanum exposed, roundish similar to half of the eye (TD/EYE 0.56), surrounded by tiny rectangular scales; ten rectangular supralabials of similar size; mental pentagonal similar in size to the adjacent infralabials; two postmentals similar in size, four scales contacting the mental; chin shield large, extending posteriorly to the angle of the jaw, separated from the infralabials by one scales row anteriorly and four at the angle of the jaw; ten rectangular infralabials, scales slightly decreasing in size posteriorly; gular sharply keeled and spinose with a yellowish, small midventral row; dewlap and gular pouch very small; nuchal crest composed of seven tiny, keeled, compressed, asymmetrical triangular spines ascending from the first to the fifth or sixth, then decreasing in height, bordered on each side by two rows of keeled, triangular scales; nuchal crest followed by a diastema of eight scales at the base of the nape; dorsal body crest is half of the nuchal crest, extending from the posterior margin of the diastema onto the sacrum; vertebral crest composed of small, epidermal, flat, saw-like triangular scales, bordered by two rows of smaller paravertebral triangular scales; vertebral crest slightly decreasing to the sacrum, then fading progressively.
Body moderately developed, narrow, triangular in cross-section; dorsal body scales small intermixed with three set of medium keeled scales line transversely from dorsal to lateral on the flanks; scales of the pectoral region and abdomen larger than the dorsal scales, keeled, semi-transverse rows arranged; keeled scales anterior to the vent large; limbs moderately long, dorsal and ventral scales of forelimbs slightly keeled, proximal scales similar to the distal scales; five digits on the manus; subdigital scales keeled, 18 subdigital lamellae under the fourth finger. Scales on the hindlimb keeled, femoral scales slightly keeled and smaller than those on the tibia; five digits on the pes; subdigital scales keeled, 21 subdigital lamellae under the fourth toe; tail length 1.16 times SVL, tail covered with keeled spinose scales, keels on subcaudals directed posteriorly; subcaudals much longer than supracaudals; base of the tail 5.3 mm wide.
The paratypes resemble the holotype in all aspects, except that females (THNHM31130 to 31134) differ from the holotype by lacking a transvers bar on the dorsal head region and possessing a pale yellowish-brown to orangish-brown body coloration. Females (THNHM31130–31134) exhibit sexual dimorphic in body size, being ~1.27 and ~1.23 times larger than males (holotype and THNHM31135 to 31136) in SVL and TAL, respectively. Morphometric and meristic data for the type series are presented in Table 2.
The hemipenis of three specimens (THNHM31129, 31135, 31136) were everted, revealing lengths of 5.3–7.2 mm. Each penis diverged into a symmetrical, sponged, budded-rosette shape with a tip width of 3.5–4.3 mm. In preserved ethanol, the hemipenis exhibited a creamy yellow coloration (Fig.
Coloration in life is characterized by a dark brown head with creamy brown transverse bars on head with the most prominent crossing the orbital region. The lips and neck region are yellowish-green for males, and whitish-brown for females. Both sexes have no black eye patch. The male body presents a dark brown marbled reticulum interspersed with some pale greenish spots, whereas the female body display large, randomly distributed whitish-yellow patterns. Whitish or whitish-yellow ocellated spots are presented at the knee and elbow, with additional marking visible on the arm and leg. The ventral coloration is creamy. Arms exhibited darker and paler marks dorsally. Legs are darker dorsally with brown bars ventrally. The male tail is banded with pale brown and a dirty yellowish-rusty coloration, while the female tail is yellowish-creamy (Figs
Molecular comparison, based on 698 nucleotides of the ND2 gene, reveal a minimal intraspecific variation among the eight analyzed specimens of A. syrax sp. nov. with pairwise sequence differences ranging from 0–0.01% (n = 8, GenBank PX738371–PX738378) (Table
Table 2. Morphometrical (in mm) and meristic data for the type series of Acanthosaura syrax sp. nov. For character abbreviations see Materials and methods. Meristic characters are given in the left column. NA = not applicable.
| Characteristics | Holotype THNHM31129 | Paratype THNHM31130 | Paratype THNHM31131 | Paratype THNHM31132 | Paratype THNHM31133 | Paratype THNHM31134 | Paratype THNHM31135 | Paratype THNHM31136 | Ranges | |
|---|---|---|---|---|---|---|---|---|---|---|
| Sex | Male | Female | Female | Female | Female | Female | Male | Male | Male | Female |
| SVL | 67.0 | 101.2 | 89.20 | 76.2 | 80.9 | 75.5 | 62.5 | 69.6 | 62.5–67.0 | 75.5–101.2 |
| TaL | 77.6 | 106.9 | 96.40 | 87.1 | 86.4 | 85.5 | 72.4 | 76.1 | 72.4–77.6 | 85.5–106.9 |
| Tal/SVL | 1.16 | 1.06 | 1.08 | 1.14 | 1.07 | 1.13 | 1.16 | 1.09 | 1.09–1.16 | 1.06–1.14 |
| TBW | 5.3 | 8.1 | 10.3 | 5.4 | 7.2 | 8.0 | 6.6 | 5.8 | 5.3–6.6 | 5.4–10.3 |
| HL | 21.1 | 30.4 | 29.7 | 23.5 | 24.4 | 24.7 | 20.5 | 20.3 | 20.3–21.1 | 23.5–30.4 |
| HL/SVL | 0.31 | 0.30 | 0.33 | 0.31 | 0.30 | 0.33 | 0.33 | 0.29 | 0.29–0.33 | 0.30–0.33 |
| HW | 14.1 | 21.2 | 21.4 | 17.5 | 16.7 | 17.7 | 15.6 | 14.4 | 14.1–15.6 | 16.7–21.4 |
| HW/SVL | 0.21 | 0.21 | 0.24 | 0.23 | 0.21 | 0.23 | 0.25 | 0.21 | 0.21–0.25 | 0.21–0.24 |
| HW/HL | 0.67 | 0.70 | 0.72 | 0.74 | 0.68 | 0.72 | 0.76 | 0.71 | 0.67–0.76 | 0.68–0.74 |
| HD | 11.1 | 17.6 | 16.8 | 12.7 | 14.4 | 14.9 | 11.8 | 10.7 | 10.7–11.8 | 12.7–17.6 |
| HD/SVL | 0.17 | 0.17 | 0.19 | 0.17 | 0.18 | 0.20 | 0.19 | 0.15 | 0.15–0.19 | 0.17–0.20 |
| SL | 7.0 | 8.0 | 9.9 | 7.8 | 7.8 | 7.9 | 6.7 | 6.7 | 6.7–7.0 | 7.8–9.9 |
| SL/SVL | 0.10 | 0.08 | 0.11 | 0.10 | 0.10 | 0.10 | 0.11 | 0.10 | 0.10–0.11 | 0.08–0.11 |
| SL/HL | 0.29 | 0.29 | 0.29 | 0.29 | 0.29 | 0.29 | 0.29 | 0.29 | 0.29 | 0.29 |
| ORBIT | 7.3 | 7.6 | 8.0 | 8.4 | 8.4 | 8.2 | 7.0 | 7.7 | 7.0–7.7 | 7.6–8.4 |
| ORB/HL | 0.11 | 0.08 | 0.09 | 0.11 | 0.10 | 0.11 | 0.11 | 0.11 | 0.11 | 0.08–0.11 |
| EYE | 4.5 | 5.2 | 5.2 | 5.2 | 5.1 | 5.3 | 5.2 | 5.0 | 4.5–5.2 | 5.1–5.3 |
| EYE/HL | 0.21 | 0.17 | 0.18 | 0.22 | 0.21 | 0.21 | 0.25 | 0.25 | 0.21–0.25 | 0.17–0.22 |
| TD | 2.5 | 3.2 | 3.1 | 2.5 | 2.2 | 2.7 | 1.9 | 2.1 | 1.9–2.5 | 2.2–3.2 |
| TD/HL | 0.16 | 0.16 | 0.16 | 0.16 | 0.16 | 0.16 | 0.16 | 0.16 | 0.16 | 0.16 |
| TD/HD | 0.23 | 0.18 | 0.18 | 0.20 | 0.15 | 0.18 | 0.16 | 0.20 | 0.16–0.23 | 0.15–0.20 |
| TN | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
| PS | 1.3 | 2.3 | 2.7 | 1.9 | 1.6 | 2.2 | 1.8 | 2.0 | 1.3–2.0 | 1.6–2.7 |
| PS/HL | 0.06 | 0.08 | 0.09 | 0.08 | 0.07 | 0.09 | 0.09 | 0.10 | 0.06–0.10 | 0.07–0.09 |
| NSL | 2.4 | 4.7 | 4.1 | 3.0 | 3.2 | 3.5 | 2.4 | 2.4 | 2.4 | 3.0–4.7 |
| NSL/HL | 0.11 | 0.15 | 0.14 | 0.13 | 0.13 | 0.14 | 0.12 | 0.12 | 0.11–0.12 | 0.13–0.15 |
| NN | 7 | 7 | 7 | 7 | 7 | 7 | 7 | 7 | 7 | 7 |
| DS | 1.5 | 2.6 | 1.4 | 1.4 | 1.4 | 1.6 | 1.4 | 1.0 | 1.0–1.5 | 1.4–2.6 |
| DS/HL | 0.07 | 0.09 | 0.05 | 0.06 | 0.06 | 0.06 | 0.07 | 0.05 | 0.05–0.07 | 0.05–0.09 |
| NSW | 0.6 | 1.1 | 1.3 | 0.5 | 1.0 | 0.7 | 0.4 | 0.5 | 0.4–0.6 | 0.5–1.3 |
| NSW/NSL | 0.25 | 0.23 | 0.32 | 0.17 | 0.31 | 0.20 | 0.17 | 0.21 | 0.17–0.25 | 0.17–0.32 |
| DIAS | 3.8 | 6.7 | 4.40 | 3.3 | 4.2 | 3.1 | 3.3 | 4.1 | 3.3–4.1 | 3.1–6.7 |
| DIAS/SVL | 0.06 | 0.07 | 0.05 | 0.04 | 0.05 | 0.04 | 0.05 | 0.06 | 0.05–0.06 | 0.04–0.07 |
| DIASN | 8 | 8 | 8 | 7 | 5 | 6 | 7 | 7 | 7–8 | 5–8 |
| FOREL | 29.1 | 44.5 | 39.50 | 35.3 | 34.4 | 31.7 | 27.9 | 33.1 | 27.9–33.1 | 31.7–44.5 |
| FOREL/SVL | 0.43 | 0.44 | 0.44 | 0.46 | 0.43 | 0.42 | 0.45 | 0.48 | 0.43–0.48 | 0.42–0.46 |
| HINDL | 33.4 | 49.3 | 45.40 | 39.8 | 38.8 | 37.3 | 33.8 | 35.9 | 33.4–35.9 | 37.3–49.3 |
| HINDL/SVL | 0.50 | 0.49 | 0.51 | 0.52 | 0.48 | 0.49 | 0.54 | 0.52 | 0.50–0.54 | 0.48–0.52 |
| FOREL/HINDL | 0.87 | 0.90 | 0.87 | 0.89 | 0.89 | 0.85 | 0.83 | 0.92 | 0.83–0.92 | 0.85–0.90 |
| SUPRAL | 10 | 11 | 10 | 9 | 10 | 10 | 11 | 10 | 10–11 | 9–11 |
| INFRAL | 10 | 11 | 10 | 10 | 11 | 10 | 10 | 10 | 10 | 10–11 |
| VENT | 60 | 58 | 57 | 60 | 59 | 56 | 56 | 58 | 56–60 | 56–60 |
| FI | 18 | 19 | 19 | 20 | 17 | 17 | 17 | 18 | 17–18 | 17–20 |
| TO | 21 | 19 | 20 | 19 | 19 | 17 | 21 | 22 | 21–22 | 17–20 |
| OS | 1.8 | 4.1 | 2.60 | 2.9 | 2.6 | 2.6 | 2.4 | 1.9 | 1.8–2.4 | 2.6–4.1 |
| PS/OS | 0.72 | 0.56 | 1.04 | 0.66 | 0.62 | 0.85 | 0.75 | 1.05 | 0.72–1.05 | 0.56–1.04 |
| OS/HL | 0.09 | 0.13 | 0.09 | 0.12 | 0.11 | 0.11 | 0.12 | 0.09 | 0.09–0.12 | 0.09–0.13 |
| NSSOS | 5 | 6 | 5 | 5 | 6 | 6 | 5 | 5 | 5 | 5–6 |
| CS | 12 | 13 | 14 | 12 | 13 | 13 | 12 | 12 | 12 | 12–14 |
| RW | 2.1 | 3.5 | 2.9 | 2.0 | 2.1 | 3.3 | 1.8 | 1.3 | 1.3–2.1 | 2.0–3.5 |
| RH | 0.9 | 1.5 | 1.1 | 1.3 | 0.8 | 1.2 | 0.7 | 0.8 | 0.7–0.9 | 0.8–1.5 |
| RS | 6 | 7 | 7 | 6 | 6 | 7 | 6 | 6 | 6 | 6–7 |
| NS | 6 | 7 | 5 | 6 | 6 | 7 | 6 | 6 | 6 | 5–7 |
| NCS | 8 | 7 | 8 | 8 | 9 | 8 | 8 | 8 | 8 | 7–9 |
| NSCSL | 7 | 7 | 7 | 7 | 8 | 6 | 7 | 6 | 6–7 | 6–8 |
| NR | 2 | 2 | 2 | 2 | 2 | 2 | 2 | 2 | 2 | 2 |
| NSSLC | 10 | 10 | 10 | 9 | 9 | 10 | 9 | 9 | 9–10 | 9–10 |
| MW | 1.1 | 1.7 | 1.0 | 1.8 | 1.0 | 1.2 | 0.7 | 1.0 | 0.7–1.1 | 1.0–1.8 |
| MH | 0.7 | 1.5 | 1.2 | 1.4 | 1.2 | 1.2 | 0.7 | 1.1 | 0.7–1.1 | 1.2–1.5 |
| MW/MH | 1.57 | 1.13 | 0.83 | 1.29 | 0.83 | 1.00 | 1.00 | 0.91 | 0.91–1.57 | 0.83–1.29 |
| PM | 4 | 4 | 4 | 4 | 4 | 4 | 4 | 4 | 4 | 4 |
| YAS | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 |
| ND | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 |
| LKP | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 |
| BEP | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
| ESBO | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
| GP | 1 | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 |
| OF | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 |
| NEBO | 9 | 10 | 9 | 10 | 10 | 9 | 9 | 10 | 9–10 | 9–10 |
| NSSOB | 4 | 4 | 5 | 5 | 4 | 4 | 4 | 4 | 4 | 4–5 |
| PL | 5.3 | NA | NA | NA | NA | NA | 7.0 | 7.2 | 5.3–7.2 | NA |
| PW | 3.8 | NA | NA | NA | NA | NA | 3.5 | 4.3 | 3.5–4.3 | NA |
| EGG | NA | 6 | NA | 16 | 11 | 6 | NA | NA | NA | 6–16 |
Furthermore, the combined Maximum–Likelihood (ML) and Bayesian Inference (BI) phylogenetic analyses of the ND2 gene strongly supported the distinct taxonomic status of A. syrax sp. nov. (76/0.78). The reconstructed phylogenies were topologically consistent, recovering the new species as a highly supported monophyletic clade completely separated from all other included taxa. Importantly, the robust nodal support specifically confirms the evolutionary divergence between A. syrax sp. nov. and its morphologically similar congener, A. lepidogaster (Fig.
Phylogram of investigated members of Acanthosaura inferred from the ND2 gene. Nodal numbers represent Maximum Likelihood bootstrap support values (BS, left) and Bayesian Inference posterior probabilities (PP, right). Values below 50% for ML or 0.70 for BI are indicated by a hyphen (-) or are not shown.
Additionally, molecular comparison, based on 600 nucleotides of the COI gene, also reveal minimal intraspecific variation among the eight analyzed specimens of A. syrax sp. nov. with pairwise sequence differences ranging from 0–0.34% (n = 8, GenBank PZ164171–PZ164178) (Table
Percentage pairwise genetic distances based on the ND2 gene of among and within nine species of the genus Acanthosaura, including the outgroup Calotes emma (see Appendix
| Taxa | 1 | 2 | 3 | 4 | 5 | 6 | 7 | 8 | 9 |
|---|---|---|---|---|---|---|---|---|---|
| Calotes emma | – | ||||||||
| Acanthosaura syrax sp. nov. | 1.91–1.94 | 0.00–0.01 | |||||||
| Acanthosaura armata | 2.17 | 0.16 | 0.00 | ||||||
| Acanthosaura aurantiacrista | 2.22–2.25 | 0.17–0.18 | 0.16 | 0.00 | |||||
| Acanthosaura capra | 2.19 | 0.17 | 0.18 | 0.19–0.20 | – | ||||
| Acanthosaura cardamomensis | 2.21–2.29 | 0.16–0.18 | 0.17–0.18 | 0.13–0.15 | 0.20–0.22 | 0.01 | |||
| Acanthosaura crucigera | 1.96–2.17 | 0.17–0.21 | 0.16–0.22 | 0.01–0.15 | 0.19–0.24 | 0.13–0.16 | 0.00–0.14 | ||
| Acanthosaura lepidogaster | 1.96–2.12 | 0.14–0.16 | 0.18–0.20 | 0.18–0.20 | 0.17–0.19 | 0.18–0.23 | 0.18–0.25 | 0.09 | |
| Acanthosaura meridiona | 2.09 | 0.20 | 0.20 | 0.12 | 0.22 | 0.15–0.16 | 0.01–0.11 | 0.21–0.24 | 0.00 |
Percentage pairwise genetic distances based on the COI gene of among and within nine species of the genus Acanthosaura, including the outgroup Calotes emma (see Appendix
| Taxa | 1 | 2 | 3 | 4 | 5 | 6 | 7 | 8 | 9 | 10 | 11 | 12 | 13 | 14 | 15 |
|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
| Calotes versicolor | – | ||||||||||||||
| Acanthosaura syrax sp. nov. | 26.83–27.09 | 0.00–0.34 | |||||||||||||
| Acanthosaura brachypoda | 28.63–29.16 | 17.30–17.75 | 0.00–0.54 | ||||||||||||
| Acanthosaura capra | 27.34 | 14.96–15.44 | 25.08–25.32 | 0.00 | |||||||||||
| Acanthosaura crucigera | 29.68 | 16.82–17.25 | 17.98–18.20 | 25.68 | 0.00 | ||||||||||
| Acanthosaura murphyi | 27.34–28.37 | 25.68–27.14 | 25.57–26.58 | 6.62–6.81 | 24.72–25.20 | 0.17–1.03 | |||||||||
| Acanthosaura coronata | 27.60–28.63 | 25.82–26.58 | 27.09–28.11 | 18.43–19.58 | 25.32–25.82 | 19.12–20.27 | 0.00–2.35 | ||||||||
| Acanthosaura cuongi | 30.74 | 26.74–27.45 | 27.09–27.60 | 17.37–17.72 | 26.00–26.46 | 18.25–18.59 | 15.75–16.42 | 0.00–0.35 | |||||||
| Acanthosaura grismeri | 29.07–29.16 | 27.64–28.13 | 29.07–29.42 | 19.30–19.44 | 27.76–27.95 | 20.56–21.46 | 8.25–8.56 | 16.42–16.85 | 0.00 | ||||||
| Acanthosaura lepidogaster | 29.68–30.74 | 16.64–17.53 | 4.01–4.77 | 24.58–25.57 | 17.08–17.98 | 24.09–25.82 | 25.57–27.09 | 26.58–27.34 | 26.73–27.09 | 0.00–1.62 | |||||
| Acanthosaura liui | 30.41–30.68 | 19.08–19.55 | 21.94–22.43 | 26.70–26.96 | 18.38–18.62 | 26.44–27.22 | 28.53–29.87 | 25.93–26.44 | 28.80–29.06 | 19.55–20.02 | 0.00–0.18 | ||||
| Acanthosaura longicaudata | 29.16 | 17.90–18.34 | 10.26 | 23.54 | 18.12 | 23.54–24.01 | 24.83–25.57 | 26.71–26.95 | 24.95–25.68 | 9.45–10.26 | 21.22–21.46 | 0.00 | |||
| Acanthosaura rubrilabris | 30.21–30.48 | 16.18–17.04 | 13.38–14.02 | 24.72–25.20 | 16.82–17.47 | 24.49–25.92 | 24.58–25.82 | 23.83–24.77 | 25.68–26.98 | 12.33–13.59 | 18.38–19.32 | 13.67–14.09 | 0.00–0.86 | ||
| Acanthosaura nataliae | 34.87–35.15 | 20.04–20.74 | 20.04–20.50 | 31.28–31.55 | 19.81–20.04 | 30.74–31.28 | 29.95–30.74 | 30.74–31.28 | 30.74–30.95 | 20.74–21.21 | 23.41–23.91 | 20.50–20.74 | 19.81–20.97 | 0.00–0.18 | |
| Acanthosaura phongdienensis | 29.95–30.74 | 15.75–16.19 | 14.88–15.10 | 25.57–26.08 | 18.20–18.66 | 25.08–26.08 | 25.08–26.08 | 27.09–27.60 | 27.60–28.28 | 13.81–14.88 | 19.79–20.50 | 12.96 | 5.15–6.10 | 22.16–22.40 | 0.00–0.54 |
These significant divergence values firmly support the species distinct taxonomic status. Concordant with the ND2 results, the dual ML and BI phylogenetic analyses of the COI dataset recovered A. syrax sp. nov. as an independent, well-supported monophyletic lineage (99/1.00) that is clearly distinguished from all other sampled Acanthosaura taxa (Fig.
Phylogram of investigated members of Acanthosaura inferred from the COI gene. Nodal numbers represent Maximum Likelihood bootstrap support values (BS, left) and Bayesian Inference posterior probabilities (PP, right). Values below 50% for ML or 0.70 for BI are indicated by a hyphen (-) or are not shown. * = 96/0.83, + = 99/0.99.
This species has a restricted altitudinal distribution within the Mae Wong National Park, where it is typically observed on the upper mountain range, exceeding 1,300 m a.s.l. (Fig.
Acanthosaura syrax sp. nov. occurs in northwestern Thailand according to personal field observations restricted to areas exceeding 1,000 m a.s.l. in Mae Wong National Park, Tak–Kamphaeng Phet Province, Thailand (16°06'01.6"N, 99°06'28.0"E).
The specific epithet syrax is a noun in apposition, named in reference to the dragon “Syrax”, the mount of Queen Rhaenyra Targaryen. This name is inspired by the species’ distinctive yellow-hued coloration, its relatively diminutive size compared to its congeners, and its significant reproductive output, reflected in its high clutch size. We suggest the following common names, “Syrax mountain horned dragon” (English) and “King Ka Khaow Naam Lek Syrax” (กิ้งก่าเขาหนามเล็กไซแรกซ์) (Thai).
Suppl. material
Acanthosaura syrax sp. nov. differs from A. armata in presenting a smaller TaL/SVL ratio (1.06–1.16 vs 1.20–1.60), smaller PS/HL ratio (0.06–0.10 vs 0.22–0.58), smaller NSL/HL ratio (0.11–0.15 vs 0.22–0.51), smaller DS/HL ratio (0.05–0.09 vs 0.20–0.52), smaller OS/HL ratio (0.09–.13 vs 0.16–0.43), fewer NS (7 vs 12), fewer INFRAL (10–11 vs 12–15), and fewer NCS (7–9 vs 10–17).
Acanthosaura syrax sp. nov. differs from A. aurantiacrista in presenting a smaller TaL/SVL ratio (1.06–1.16 vs 1.40–1.70), smaller SL/HL (0.29 vs 0.41–.58), smaller PS/HL ratio (0.06–0.10 vs 0.24–0.84), smaller NSL/HL ratio (0.11–0.15 vs 0.35–0.95), smaller DS/HL ratio (0.05–0.09 vs 0.15–0.38), greater NSW/NSL (0.17–.32 vs 0.07–0.14), smaller OS/HL ratio (0.09–.13 vs 0.19–0.44), greater FOREL/HINDL (0.83–0.92 vs 0.75–0.80), fewer VENT (56–60 vs 63–66), fewer TO (17–22 vs 25–29), smaller PS/OS (0.56–1.05–1.08–1.91), fewer NS (7 vs 8), and fewer NCS (7–9 vs 11–13).
Acanthosaura syrax sp. nov. differs from A. cardamomensis in presenting a smaller SL/HL (0.29 vs 0.47–0.57), smaller PS/HL ratio (0.06–0.10 vs 0.14–0.45), smaller NSL/HL ratio (0.11–0.15 vs 0.17–0.66), smaller DS/HL ratio (0.05–0.09 vs 0.14–0.45), smaller NSW (0.4–1.3 vs 1.8–4.2), smaller OS/HL ratio (0.09–0.13 vs 0.24–0.56), slightly fewer NS (5–7 vs 7–10), slightly fewer NCS (7–9 vs 9–17), slightly fewer NSSLC (9–10 vs 10–19), and the absence of black eye patch.
Acanthosaura syrax sp. nov. differs from A. crucigera in presenting of a smaller ORBIT (7.0–8.4 vs 8.9–10.8), slightly smaller PS/HL ratio (0.06–0.10 vs 0.09–0.33), slightly smaller DS/HL ratio (0.05–0.09 vs 0.09–0.24), slightly smaller NSW (0.4–1.3 vs 1.3–3.4), fewer DIASN (5–8 vs 9–25), slightly fewer NCS (7–9 vs 9–12), slightly fewer NSSLC (9–10 vs 10–14), and the absence of black eye patch.
Acanthosaura syrax sp. nov. is most closely related to A. lepidogaster. Acanthosaura syrax sp. nov. differs from A. lepidogaster in presenting a smaller TaL/SVL ratio (1.06–1.16 vs 1.6–1.9), greater HD (14.9–17.6 vs 12.0–12.5), fewer DIASN (5–8 vs 10–14), slightly fewer TO (17–22 vs 22–23), greater OS/HL ratio (0.09–0.13 vs 0.14–0.15), and fewer PM (4 vs 5).
Acanthosaura syrax sp. nov. differs from A. meridiona in presenting a smaller TaL (72.4–106.9 vs 108.5–176.0), greater HL (24.7–30.4 vs 17.0–24.1), smaller ORBIT/HL ratio (0.08–0.11 vs 0.44–0.53), smaller PS/HL ratio (0.06–0.10 vs 0.16–0.38), slightly smaller NSL/HL ratio (0.11–0.15 vs 0.15–0.37), fewer DIASN (5–8 vs 10–16), slightly fewer VENT (56–60 vs 60–68), slightly smaller OS/HL ratio (0.09–0.13 vs 0.13–0.25), fewer NCS (7–9 vs 10–15), fewer NSCSL (6–8 vs 9–11), and more NR (2 vs 1).
Acanthosaura syrax sp. nov. differs from A. phuketensis in presenting a smaller TaL/SVL ratio (1.06–1.16 vs 1.40–1.70), smaller TD (1.9–3.2 vs 3.5–4.7), smaller PS/HL (0.06–0.10 vs 0.23–0.38), smaller NSL/HL ratio (0.11–0.15 vs 0.21–0.39), smaller DS/HL ratio (0.05–0.09 vs 0.11–0.26), smaller NSW (0.4–1.3 vs 1.4–2.9), fewer DIASN (5–8 vs 12–17), fewer NCS (7–9 vs 12–13), fewer NSSLC (9–10 vs 11–14), and the absence of black eye patch.
The results indicate that A. crucigera and A. lepidogaster mostly share characteristics with A. syrax sp. nov. However, a close examination led us to separate A. syrax sp. nov. from these similar species based on coloration, morphological characteristics, especially nuchal–dorsal crest patterns, as well as on molecular data (Fig.
For other congeners outside Thailand, they are compared based on published data by
Acanthosaura syrax sp. nov. differs from A. bintangensis Wood, Grismer, Grismer, Northayati, Chan & Bauer, 2009 in presenting a smaller TaL/SVL ratio (1.06–1.16 vs 1.30–1.40), smaller NSL/HL ratio (0.11–0.15 vs 0.17–0.21), smaller NSW (0.4–1.3 vs 1.6–2.1), fewer DIASN (5–8 vs 11–15), fewer SUPRAL (9–11 vs 12), more VENT (56–60 vs 51–55), fewer FI (17–20 vs 23), fewer TO (17–22 vs 26–28), smaller RW (1.3–3.5 vs 3.6–5.3), smaller RH (0.7–1.5 vs 1.7–20), fewer NS (5–7 vs 8), fewer NCS (7–9 vs 10–11), more NR (2 vs 1), smaller GP (0–1 vs 3–4), the presence of LKP which is absent in from A. bintangensis, and the absence of black eye patch and elliptical scales below the orbit.
Acanthosaura syrax sp. nov. differs from A. brachypoda in presenting a smaller TaL/SVL ratio (1.30–1.40 vs 1.58), smaller SL (6.7–9.9 vs 12.2), smaller TD (1.9–3.2 vs 3.6), smaller PS/HL ratio (0.06–0.10 vs 0.11), smaller NSL/HL ratio (0.11–0.15 vs 0.16), smaller NSW (0.4–1.3 vs 1.6), fewer SUPRAL (9–11 vs 12–13), fewer VENT (56–60 vs 63), fewer TO (17–22 vs 24), greater OS/HL ratio (0.09–0.13 vs 0.03), smaller RH (0.7–1.5 vs 2.3), fewer NS (5–7 vs 9), smaller MW (0.7–1.8 vs 2.9), smaller MH (0.7–1.5 vs 2.1), and the absence of black eye patch.
Acanthosaura syrax sp. nov. differs from A. capra in presenting a smaller TaL/SVL ratio (1.06–1.16 vs 1.20–1.50), smaller TD (1.9–3.2 vs 3.4–5.2), smaller PS/HL ratio (0.06–0.10 vs 0.36), smaller NSL/HL ratio (0.11–0.15 vs 0.42–0.43), smaller DS/HL ratio (0.05–0.09 vs 0.16–0.17), smaller FOREL (27.9–44.5 vs 54.2–83.8), smaller HINDL (33.4–49.3 vs 78.5–107.2), fewer INFRAL (10–11 vs 12–13), smaller RW (1.3–3.5 vs 4.2–4.6), smaller RH (0.7–1.5 vs 1.8–2.3), fewer NS (5–7 vs 9), smaller MW (0.7–1.8 vs 1.9–2.2), smaller GP (0–1 vs 3–4), the presence of OS, NSSOS which is absent in A. capra, and the absence of black eye patch.
Acanthosaura syrax sp. nov. differs from A. coronata in presenting a greater TaL/SVL ratio (1.06–1.16 vs 0.6–1.0), greater HL (24.7–30.4 vs 14.4–16.3), greater HW (17.7–21.2 vs 13.6–17.5), fewer SUPRAL (9–11 vs 12–13), more FI (17–20 vs 13–14), greater RW (1.3–3.5 vs 0.8–0.9), fewer RS (6–7 vs 9), fewer NR (2 vs 3–4), and the presence of postorbital spines, nuchal scales, dorsal scales, diastema, occipital spines, and black, diamond shaped, nuchal collar, which is absent in A. coronata.
Acanthosaura syrax sp. nov. differs from A. cuongi in presenting a smaller TaL (72.4–106.9 vs 113.9–132.6), slightly greater HW/SVL (0.21–0.25 vs 0.18–0.21), smaller ORBIT/HL (0.08–0.11 vs 0.27–0.38), smaller EYE (4.5–5.3 vs 6.2–7.5), slightly greater PS/HL (0.06–0.10 vs 0.00–0.06), greater NSL/HL (0.11–0.15 vs 0.06–0.10), smaller NSW/NSL (0.17–0.32 vs 0.41–0.76), fewer NS (7 vs 8), greater FOREL/SVL (0.42–0.48 vs 0.31–0.38), greater FOREL/HINDL (0.83–0.92 vs 0.70–0.78), slightly more FI (17–20 vs 14–17), greater OS/HL (0.09–0.13 vs 0.04–0.08), greater CS (12–14 vs 10–11), fewer NS (5–7 vs 8–10), greater NR (2 vs 1), and the absence of black eye patch.
Acanthosaura syrax sp. nov. differs from A. grismeri in presenting a smaller TaL/SVL (72.4–106.9 vs 1.21–1.44), TBW (5.3–10.3 vs 11.1–13.0), SL/HL (0.29 vs 0.31–0.40), ORBIT/HL (0.08–0.11 vs 0.32–0.43), TD (1.9–3.2 vs 3.2–5.3), NSW/NSL (0.17–0.32 vs 0.60–1.04), OS/HL (0.09–0.13 vs 0.40–1.94), greater HW/SVL (0.21–0.25 vs 0.19–0.20), DS/HL (0.05–0.09 vs 0.03–0.04), FOREL/SVL (0.42–0.48 vs 0.29–0.34), FOREL/HINDL (0.83–0.92 vs 0.66–0.77), FI (17–20 vs 12–16), PS/OS (0.56–1.05 vs 0.04–0.09), RS (6–7 vs 4–5), the presence diastema and oblique fold.
Acanthosaura syrax sp. nov. differs from A. liui in presenting a smaller TaL/SVL ratio (1.06–1.16 vs 1.47–1.77), smaller TBW (5.3–10.3 vs 10.6–13.9), smaller SL (6.7–9.9 vs 10.3–11.3), slightly smaller ORBIT (7.0–8.4 vs 8.4–8.9), smaller EYE (4.5–5.3 vs 5.8–6.4), greater TD/HD ratio (0.15–0.23 vs 0.03–0.04), smaller NSL/HL ratio (0.11–0.15 vs 0.17–0.24), smaller DS/HL ratio (0.05–0.09 vs 0.10–0.12), smaller HINDL (33.4–49.3 vs 50.7–52.6), slightly fewer TO (17–22 vs 22–25), fewer RS (6–7 vs 8–9), smaller MH (0.7–1.5 vs 1.7–2.3), fewer PM (4 vs 5–6), smaller GP (0–1 vs 2–3), and the absence of black eye patch.
Acanthosaura syrax sp. nov. differs from A. longicaudata in presenting a smaller TaL/SVL ratio (1.06–1.16 vs 1.62–2.04), greater HW (17.7–21.2 vs 14.2–17.4), smaller SL/HL ratio (0.29 vs 0.35–0.37), fewer ORBIT/HL (0.08–0.11 vs 0.32–0.37), slightly greater TD (1.9–3.2 vs 1.4–1.9), smaller NSW/NSL ratio (0.17–0.32 vs 0.46–0.64), HINDL/SVL ratio (0.48–0.54 vs 0.55–0.63), FOREL/HINDL ratio (0.83–0.92 vs 0.64–0.75), and slightly more VENT (56–60 vs 47–56).
Acanthosaura syrax sp. nov. differs from A. murphyi in presenting a smaller TaL/SVL ratio (1.06–1.16 vs 1.48–1.54), smaller HD (14.9–17.6 vs 18.5–20.6), smaller SL (6.7–9.9 vs 10.3–15.3), smaller ORBIT (7.0–8.4 vs 9.9–12.3), smaller PS/HL ratio (0.06–0.10 vs 0.16–0.34), smaller NSL/HL ratio (0.11–0.15 vs 0.24–0.43), smaller DS/HL ratio (0.05–0.09 vs 0.14–0.51), smaller NSW (0.4–1.3 vs 2.9–4.8), smaller FOREL (27.9–44.5 vs 49.8–56.6), smaller HINDL (33.4–49.3 vs 60.4–68.4), fewer SUPRAL (9–11 vs 12–14), fewer INFRAL (10–11 vs 12–14), fewer RS (6–7 vs 8–9), fewer NCS (7–9 vs 13–16), fewer NR (2 vs 3–4), smaller GP (0–1 vs 4), the absence of tympanum scale, and the presence of occipital spines, which is absent in A. murphyi.
Acanthosaura syrax sp. nov. differs from A. nataliae in presenting a smaller SVL (62.5–101.2 vs 106.7–158.0), smaller TaL (72.4–106.9 vs 132.5–190.0), smaller SL (6.7–9.9 vs 12.0–19.9), smaller TD (1.9–3.2 vs 3.9–7.0), smaller PS/HL ratio (0.06–0.10 vs 0.36–0.52), smaller NSL/HL ratio (0.11–0.15 vs 0.58–0.75), smaller DS/HL ratio (0.05–0.09 vs 0.41–0.53), smaller FOREL (27.9–44.5 vs 58.4–85.0), smaller HINDL (33.4–49.3 vs 72.1–129.7), fewer VENT (56–60 vs 64–71), smaller RW (1.3–3.5 vs 4.6–6.1), smaller RH (0.7–1.5 vs 1.8–2.9), fewer NCS (7–9 vs 10–14), more NR (2 vs 1), fewer NSSLC (9–10 vs 13–16), smaller MW (0.7–1.8 vs 2.3–2.9), smaller MH (0.7–1.5 vs 2.0–2.9), smaller GP (0–1 vs 3–4), the absence of occipital spine, and the presence of black, diamond shaped, nuchal collar, and pale knee patches, which is absent in A. nataliae.
Acanthosaura syrax sp. nov. differs from A. phongdienensis in presenting a smaller TaL/SVL ratio (1.06–1.16 vs 1.50–1.90), greater HL (24.7–30.4 vs 18.6–23.8), greater HD (14.9–17.6 vs 10.4–13.6), and the presence of diastema, which is absent in A. phongdienensis.
Acanthosaura syrax sp. nov. differs from A. prasina Ananjeva, Ermakov, Nguyen, Nguyen, Murphy, Lukonina & Orlov, 2020 in presenting a smaller TaL/SVL ratio (1.06–1.16 vs 1.64–20.70), greater HL (24.7–30.4 vs 20.7–22.4), greater HW/HL ratio (0.21–0.25 vs 0.17–0.20), greater HW (14.9–17.6 vs 12.3–13.3), smaller ORBIT/HL ratio (0.08–0.11 vs 0.31–0.39), fewer TO (17–22 vs 23–26), and more CS (12–14 vs 5–6).
Acanthosaura syrax sp. nov. differs from A. rubrilabris in presenting a smaller TaL/SVL ratio (1.06–1.16 vs 1.31–1.88), smaller ORBIT/HL ratio (0.08–0.11 vs 0.28–0.35), greater TD/HL (0.16 vs 0.08–0.15), greater NS (7 vs 3–6), smaller NSW/NSL ratio (0.17–0.32 vs 0.43–0.89), greater FOREL/HINDL ratio (0.83–0.92 vs 0.67–0.78), and the absence of black eye patch.
Acanthosaura syrax sp. nov. differs from A. titiwangsaensis in presenting a smaller TaL (72.4–106.9 vs 136.0–174.0), greater HL (24.7–30.4 vs 20.0–24.3), smaller PS/HL ratio (0.06–0.10 vs 0.14–0.18), smaller NSW (0.4–1.3 vs 1.4–1.6), fewer DIASN (5–8 vs 10–13), fewer SUPRAL (9–11 vs 12–13), fewer TO (17–22 vs 23–27), smaller RW (1.3–3.5 vs 3.6–5.2), more RS (6–7 vs 5), fewer NS (5–7 vs 8), fewer NCS (7–9 vs 11–12), fewer NSCSL (6–8 vs 9–11), fewer NSSLC (9–10 vs 11–14), fewer PM (4 vs 5), smaller GP (0–1 vs 2–4), the absence of black eye patch, and the presence of pale knee patches, which is absent in A. titiwangsaensis.
Acanthosaura syrax sp. nov. differs from A. tongbiguanensis Liu & Rao, 2019 in presenting a smaller TaL/SVL ratio (1.06–1.16 vs 1.56–1.85), slightly smaller TD (1.9–3.2 vs 3.2–4.2), smaller PS/HL ratio (0.06–0.10 vs 0.13–0.19), slightly smaller NSL/HL ratio (0.11–0.15 vs 0.15–0.21), slightly smaller DS/HL ratio (0.05–0.09 vs 0.09–0.13), smaller HINDL (33.4–49.3 vs 54.1–63.9), fewer TO (17–22 vs 25–28), smaller OS/HL ratio (0.09–0.13 vs0.16–0.23), fewer NS (5–7 vs 8–9), fewer NCS (7–9 vs 10–13), and the absence of black eye patch.
The discovery of Acanthosaura syrax sp. nov. indicates that the species diversity of the genus Acanthosaura in Thailand has been significantly underestimated for a time. While most members of this genus have been described recently in other countries in Southeast Asia, particularly China and Vietnam, the ongoing discovery of new species, including this 23rd recognized taxon, suggests that high-elevation mountain ranges still harbor substantial undocumented biodiversity (
Morphologically, the unique characteristics of A. syrax sp. nov. further justify its distinct status. It is characterized by its small body size (SVL 62.4–78.5 mm) and possessing the smallest nuchal and dorsal scales recorded in the genus (NSL 2.4–4.7 mm, DS 1.0–2.6 mm). These traits suggest that A. syrax sp. nov. has evolved as a specialized high-altitude lineage, restricted primarily to the evergreen forests of the Mae Wong National Park Range at elevations above 1,300 m a.s.l.
The multi-locus molecular data generated in this study provides robust evidence supporting the status of A. syrax sp. nov. as a distinct evolutionary lineage. Analysis of the mitochondrial ND2 gene demonstrates that A. syrax sp. nov. exhibits high interspecific genetic divergence (0.14–0.21%) from all other analyzed Thai congeners. Most notably, a substantial genetic distance (0.14–0.16%) separates the new species from A. lepidogaster, its closest and morphologically most similar congener in western and northern Thailand. This high level of divergence is fully corroborated by the COI dataset, which shows deep interspecific distances between A. syrax sp. nov. and A. lepidogaster (16.64–17.53%). Furthermore, the new species maintains significant genetic isolation from other distributed congeners across China, Myanmar, and Vietnam, with COI pairwise distances ranging from 14.96% in A. capra to 28.13% in A. grismeri.
Beyond simple genetic distance thresholds, the robustness of our taxonomic conclusions is strongly reinforced by our updated phylogenetic framework. Both the Maximum Likelihood (ML) and Bayesian Inference (BI) analyses across both independent markers yielded congruent topologies (Figs
The distribution of this species appears limited to high-altitude refugia, which are increasingly vulnerable to environmental shifts. While our field surveys indicate that populations in certain areas of Mae Wong National Park remain present, they are naturally fragmented by the rugged topography of the mountain range. This habitat fragmentation poses a significant long-term threat, as isolated populations may face difficulties in gene exchange, potentially leading to inbreeding and reduced genetic fitness. Such factors are detrimental to the survival of high-altitude specialists that cannot easily migrate to lower, warmer elevations. Despite our inability to locate A. lepidogaster within this specific high-altitude area, we encountered the species approximately 50 km to the north in Khlong Lan National Park. There, A. lepidogaster occurs in low-to-mid-elevation habitats characterized by significantly warmer conditions.
Surprisingly, Acanthosaura syrax sp. nov. shares a superficial resemblance to A. lamnidentata Boulenger, 1885 (currently a synonym of A. lepidogaster) from Pegu, Myanmar, specifically in the presence of small spines and a similar body plan. However, their morphological characteristics are clearly distinct: A. syrax sp. nov. exhibits significantly smaller proportions across all key diagnostic metrics when compared to A. lamnidentata, including SVL (62.5–101.2 vs 110), TAL (72.4–106.9 vs 170), FOREL (27.90–44.50 vs 57), and HINDL (33.40–49.30 vs 92) (Boulenger, 1885). Furthermore, most morphological characteristics of A. lamnidentata align more closely with the A. lepidogaster complex than with A. syrax sp. nov. Geographically, these populations are isolated by the formidable barrier of the Tenasserim Mountains, as well as by distinct ecological conditions associated with high-elevation habitats. These significant morphological and biogeographical differences provide strong evidence that these two populations represent distinct taxa.
Consequently, we suggest that the local government and National Park authorities strengthen the protection of these high-elevation habitats. Protecting the primary evergreen forest is the most effective strategy for ensuring the survival of A. syrax sp. nov. and other sympatric fauna. We recommend adopting this new species together with other threatened creatures [e.g. Asian forest tortoise (Manouria emys Schlegel & Müller, 1844), Oldham’s leaf turtle (Cyclemys oldhamii Gray, 1863), grey-sided thrush (Turdus feae (Salvadori, 1887)), rufous-necked hornbill (Aceros nipalensis (Hodgson, 1829)), and the Indochinese tiger (Panthera tigris (Linnaeus, 1758))] as flagship representatives for the Mae Wong region to draw scientific and public attention to the conservation of Thailand’s rapidly changing mountain ecosystems. Future studies should continue to explore neighboring mountain ranges to determine the full extent of this species’ distribution and identify further cryptic diversity that likely remains hidden within the northern and western highlands.
Special thanks are due to Dr. Vet. Savek Kiatsomphob from the Department of Animal Science, Faculty of Agriculture, Kasetsart University, for his invaluable support and facilitation in obtaining ethical approval from the Institute of Animal Care and Use Committee at Kasetsart University under number ACKU68–AGR–021. We also thank to Saa-ard Thupkhunthod and Raksanan Apinunnanon from Mae Wong National Park for their kind supporting and facilitating our survey of the Mae Wong National Park.
| Taxon | Voucher | Locality | GenBank | Reference | |
|---|---|---|---|---|---|
| ND2 | COI | ||||
| Ingroup | |||||
| Acanthosaura syrax sp. nov. | THNHM31129 | Mae Wong, Kampheang Phet, Thailand | PX738375 | PZ164171 | This study |
| Acanthosaura syrax sp. nov. | THNHM31130 | Mae Wong, Kampheang Phet, Thailand | PX738377 | PZ164172 | This study |
| Acanthosaura syrax sp. nov. | THNHM31131 | Mae Wong, Kampheang Phet, Thailand | PX738378 | PZ164173 | This study |
| Acanthosaura syrax sp. nov. | THNHM31132 | Mae Wong, Kampheang Phet, Thailand | PX738371 | PZ164174 | This study |
| Acanthosaura syrax sp. nov. | THNHM31133 | Mae Wong, Kampheang Phet, Thailand | PX738372 | PZ164175 | This study |
| Acanthosaura syrax sp. nov. | THNHM31134 | Mae Wong, Kampheang Phet, Thailand | PX738373 | PZ164176 | This study |
| Acanthosaura syrax sp. nov. | THNHM31135 | Mae Wong, Kampheang Phet, Thailand | PX738374 | PZ164177 | This study |
| Acanthosaura syrax sp. nov. | THNHM31136 | Mae Wong, Kampheang Phet, Thailand | PX738376 | PZ164178 | This study |
| Acanthosaura armata | NSMT–H4595 | Asia | AB266452 | – | |
| Acanthosaura armata | – | Asia | NC014175 | – | |
| Acanthosaura aurantiacrista | THNHM28064 | Mae Sariang, Mae Hong Son, Thailand | MH777406 | – | |
| Acanthosaura aurantiacrista | QSMI1446 | Sop Khong, Omkoi, Chiang Mai, Thailand | MK798128 | – | |
| Acanthosaura aurantiacrista | THNHM28521 | Sop Khong, Omkoi, Chiang Mai, Thailand | MK798129 | – | |
| Acanthosaura aurantiacrista | THNHM28522 | Sop Khong, Omkoi, Chiang Mai, Thailand | MK798130 | – | |
| Acanthosaura aurantiacrista | QSMI1447 | Sop Khong, Omkoi, Chiang Mai, Thailand | MK798131 | – | |
| Acanthosaura aurantiacrista | QSMI1448 | Sop Khong, Omkoi, Chiang Mai, Thailand | MK798132 | – | |
| Acanthosaura aurantiacrista | THNHM28523 | Sop Khong, Omkoi, Chiang Mai, Thailand | MK798133 | – | |
| Acanthosaura brachypoda | ROM 38118 | Sa Pa, Lao Cai, Vietnam | – | MK695182 | |
| Acanthosaura brachypoda | ROM 38119 | Sa Pa, Lao Cai, Vietnam | – | MK695183 | |
| Acanthosaura brachypoda | ROM 38120 | Sa Pa, Lao Cai, Vietnam | – | MK695184 | |
| Acanthosaura capra | MVZ222130 | Vietnam | AF128498 | – | |
| Acanthosaura capra | BGM 01 | Bu Gia Map NP, Binh Phuoc (now Dong Nai), Vietnam | – | MK239022 | |
| Acanthosaura cardamomensis | FMNH263225 | Kampot, Cambodia | GU817397 | – | |
| Acanthosaura cardamomensis | FMNH263261 | Kampot, Cambodia | GU817400 | – | |
| Acanthosaura coronata | KIZ47 | Bu Gia Map NP, Binh Phuoc (now Dong Nai), Vietnam | – | MK695186 | |
| Acanthosaura coronata | KIZ48 | Bu Gia Map NP, Binh Phuoc (now Dong Nai), Vietnam | – | MK695187 | |
| Acanthosaura coronata | KIZ49 | Bu Gia Map NP, Binh Phuoc (now Dong Nai), Vietnam | – | MK695188 | |
| Acanthosaura coronata | ROM 42240 | Cat Tien NP, Dong Nai, Vietnam | – | MK695189 | |
| Acanthosaura coronata | ROM 42241 | Cat Tien NP, Dong Nai, Vietnam | – | MK695190 | |
| Acanthosaura crucigera | QSMI1592 | Muang, Tak, Thailand | MH777408 | – | |
| Acanthosaura crucigera | QSMI1593 | Muang, Tak, Thailand | MH777403 | – | |
| Acanthosaura crucigera | THNHM28057 | Muang, Tak, Thailand | MH777402 | – | |
| Acanthosaura crucigera | CAS229582 | Kawthaung, Tanintharyi, Myanmar | GU817389 | – | |
| Acanthosaura crucigera | CUMZR2008.05.26.1 | Petchaburi, Thailand | HM143889 | – | |
| Acanthosaura crucigera | USNM587487 | Tanintharyi, Ma Noe Lone, Lenya, Myanmar | – | MT607982 | |
| Acanthosaura crucigera | USNM587488 | Tanintharyi, Ma Noe Lone, Lenya, Myanmar | – | MT607980 | |
| Acanthosaura crucigera | USNM587489 | Tanintharyi, Ma Noe Lone, Lenya, Myanmar | – | MT607981 | |
| Acanthosaura cuongi | IEBR R.5250 | Phu Yen (now Dak Lak), Vietnam | – | PP669986 | |
| Acanthosaura cuongi | IEBR R.5257 | Phu Yen (now Dak Lak), Vietnam | – | PP669987 | |
| Acanthosaura cuongi | IEBR R.5251 | Khanh Hoa, Vietnam | – | PP669985 | |
| Acanthosaura grismeri | IEBR-R6353 | Krong Pong forest, Dak Lak, Vietnam | – | PV646694 | |
| Acanthosaura grismeri | IEBR-R6354 | Krong Pong forest, Dak Lak, Vietnam | – | PV646695 | |
| Acanthosaura grismeri | IEBR-R6357 | Krong Pong forest, Dak Lak, Vietnam | – | PV646696 | |
| Acanthosaura lepidogaster | MVZ224090 | Vinh Thu, Vietnam | AF128499 | – | |
| Acanthosaura lepidogaster | MD001 | Hainan, China | KR092427 | – | |
| Acanthosaura lepidogaster | ROM 30505 | Na Hang NR, Tuyen Quang, Vietnam | – | MK695191 | |
| Acanthosaura lepidogaster | ROM 30507 | Na Hang NR, Tuyen Quang, Vietnam | – | MK695192 | |
| Acanthosaura lepidogaster | ROM 30677 | Quang Thanh, Cao Bang, Vietnam | – | MK695193 | |
| Acanthosaura lepidogaster | ROM 30680 | Na Hang NR, Tuyen Quang, Vietnam | – | MK695194 | |
| Acanthosaura lepidogaster | ROM 30712 | Tam Dao NP, Vinh Phuc (now Phu Tho), Vietnam | – | MK695195 | |
| Acanthosaura liui | KIZL2020003 | Jianshui, Honghe, Yunnan, China | – | MT769763 | |
| Acanthosaura liui | KIZL2020004 | Jianshui, Honghe, Yunnan, China | – | MT769764 | |
| Acanthosaura liui | KIZL2020005 | Jianshui, Honghe, Yunnan, China | – | MT769765 | |
| Acanthosaura liui | KIZL2020006 | Jianshui, Honghe, Yunnan, China | – | MT769766 | |
| Acanthosaura liui | KIZL2020007 | Jianshui, Honghe, Yunnan, China | – | MT769767 | |
| Acanthosaura longicaudata | KIZ_L0051 | Jiangcheng, Yunnan, China | – | ON207528 | |
| Acanthosaura longicaudata | KIZ_L0132 | Jiangcheng, Yunnan, China | – | ON207529 | |
| Acanthosaura longicaudata | KIZ_L0133 | Jiangcheng, Yunnan, China | – | ON207530 | |
| Acanthosaura longicaudata | KIZ_L0150 | Jiangcheng, Yunnan, China | – | ON207531 | |
| Acanthosaura meridiona | QSMI1594 | Na Yong, Trang, Thaiand | MH777404 | – | |
| Acanthosaura meridiona | THNHM28061 | Na Yong, Trang, Thaiand | MH777407 | – | |
| Acanthosaura meridiona | THNHM28062 | Na Yong, Trang, Thaiand | MH777405 | – | |
| Acanthosaura murphyi | ITBCZ 4603 | Deo Ca Forest, Phu Yen (now Dak Lak), Vietnam | – | MK239025 | |
| Acanthosaura murphyi | ITBCZ 3533 | Hon Ba NR, Khanh Hoa, Vietnam | – | MK239026 | |
| Acanthosaura nataliae | ROM30629 | Tram Lap, Gia Lai, Vietnam | – | MK695202 | |
| Acanthosaura nataliae | ROM30631 | Tram Lap, Gia Lai, Vietnam | – | MK695203 | |
| Acanthosaura nataliae | ROM30632 | Tram Lap, Gia Lai, Vietnam | – | MK695204 | |
| Acanthosaura phongdienensis | ITBCZ6831 | Phong Dien, Thua Thien Hue, Vietnam | – | MK695205 | |
| Acanthosaura phongdienensis | ROM48013 | Phong Dien, Thua Thien Hue, Vietnam | – | MK695206 | |
| Acanthosaura phongdienensis | KIZ10657 | Phong Dien, Thua Thien Hue, Vietnam | – | MK695207 | |
| Acanthosaura phongdienensis | ITBCZ6830 | Phong Dien, Thua Thien Hue, Vietnam | – | MK695208 | |
| Acanthosaura phongdienensis | KIZ10695 | Phong Dien, Thua Thien Hue, Vietnam | – | MK695209 | |
| Acanthosaura phongdienensis | KIZ10697 | Phong Dien, Thua Thien Hue, Vietnam | – | MK695210 | |
| Acanthosaura rubrilabris | KIZ_L0057 | Menghai, Yunnan, China | – | ON207534 | |
| Acanthosaura rubrilabris | KIZ_L0010 | Menghai, Yunnan, China | – | ON207535 | |
| Acanthosaura rubrilabris | KIZ_L0006 | Jinghong, Yunnan, China | – | ON207536 | |
| Acanthosaura rubrilabris | KIZ_L0040 | Jinghong, Yunnan, China | – | ON207537 | |
| Acanthosaura rubrilabris | KIZ_L0137 | Lancang, Yunnan, China | – | ON207543 | |
| Acanthosaura rubrilabris | KIZ_L0140 | Lancang, Yunnan, China | – | ON207544 | |
| Outgroup | |||||
| Calotes emma | CAS223062 | Rakhine State, Myanmar | DQ289460 | – | |
| Calotes emma | KIZ1120 | Ta Kou NR, Binh Thuan (now Lam Dong), VN | – | MK695211 | |
The authors have declared that no competing interests exist.
This research is approved for animal ethics from the Institute of Animal Care and Use Committee at Kasetsart University under number ACKU68–AGR–021.
The authors accept full responsibility for the content of the manuscript, including the disclosure of any use of AI.
No AI tools were used in the preparation of this manuscript.
This research project was approved and funded by the Chulabhorn Royal Academy (E2567–074). All essential tools and facilities were supported by the Department of Zoology, Faculty of Science, Kasetsart University and the Panalai Veterinary Hospital.
Poramad Trivalairat: field survey, specimen examination, photograph, measurement, DNA technique, manuscript drafting. Krittiya Trivalairat: field survey, specimen examination, manuscript drafting. Nawee Moungkhaek: document management, specimen examination, measurement. Simon Caponov: field survey, local communication, specimen examination. Benchapol Lorsunyaluck: specimen examination, photograph, dissection.
P. Trivalairat https://orcid.org/0000-0001-8849-8517
K. Trivalairat https://orcid.org/0000-0003-0005-0801
B. Lorsunyaluck https://orcid.org/0000-0002-0300-5937
All of the data that support the findings of this study are available in the main text or Supplementary Information.
Supplementary tables
Data type: xlsx
Explanation note: table SS1. The comparisons of the morphometric measurements and meristic data for all currently recognised species in comparison with A. syrax sp. nov. and other recognised Acanthosaura species. table S2. Morphological comparison of nuchal and dorsal crest heights among Acanthosaura species.